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Home , Alcyonacea, Alcyoniidae, Anthelia (coral), Briareidae, Briareum, Capnella

Zoological Studies 43(3): 548-560 (2004)

Soft (: ) from Southern Taiwan. II. Diversity and Distributional Patterns Yehuda Benayahu1,*, Ming-Shiou Jeng2, Shimrit Perkol-Finkel1 and Chang-Feng Dai3 1Department of Zoology, George S. Wise Faculty of Life Sciences, Tel Aviv University, Ramat Aviv, Tel Aviv 69978, Israel 2Institute of Zoology, Academia Sinica, Taipei, Taiwan 115, R.O.C. 3Institute of Oceanography, National Taiwan University, Taipei, P.O. Box 23-13, Taiwan 106, R.O.C.

(Accepted June 28, 2004)

Yehuda Benayahu, Ming-Shiou Jeng, Shimrit Perkol-Finkel and Chang-Feng Dai (2004) Soft corals (Octocorallia: Alcyonacea) from southern Taiwan. II. species diversity and distributional patterns. Zoological Studies 43(3): 548-560. The geographic setting of Taiwan, between the West Pacific Ocean and East China Sea and at the crossroads of the Philippine-Japan I. Arc, has produced reefs of special biogeographical inter- est. It has been suggested that Taiwan may have served as a“stepping-stone”for the dispersal of shallow reef organisms. Soft corals from Nanwan Bay and Green I. (Lutao in Chinese), southern Taiwan, were studied dur- ing 1994 and 1998. Scuba collections were carried out to a depth of 33 m. Approximately 230 samples were collected, encompassing a variety of species found on these reefs. Visual estimates were made of the under- water abundance of species. The collection yielded 69 species of the families Helioporidae, , Tubiporidae, , , , and . These included 1 new species and 43 new zoogeographical records for Taiwan. Among the 22 listed genera, the survey recorded 7 for the 1st time in Taiwanese reefs. The findings confirm the high soft- diversity of these reefs. The shallow reefs of Nanwan Bay are densely inhabited by species of the family Alcyoniidae. In contrast, the deep reefs are characterized only by sporadic colonies of Alcyoniidae, but abundant assemblages of azooxanthellate members of the fami- lies Nephtheidae and . Soft corals of the family Xeniidae are abundant on the reefs of Green I. at a depth range of 3~10 m. Interestingly, species of the family Xeniidae were rarely observed at Nanwan Bay. The coral reefs of Taiwan and Japan are closely linked by the northward-flowing Kuroshio Current, which brings water and larvae from the reefs of the South China Sea. Therefore, we compared the generic affiliation and abundance estimates of the Alcyoniidae between southern Taiwan and the Ryukyu Archipelago (Japan), and found a close resemblance between these 2 reef areas. http://www.sinica.edu.tw/zool/zoolstud/43.3/548.pdf

Key words: Octocorallia, Taiwan, Coral reefs, Species diversity, Biogeography.

Taiwan, a continental island with several off- crossroads of the Philippine-Japan I. Arc, has pro- shore islets, has coral reefs that sit at the northern duced reefs of special biogeographical interest. edge of the Southeast Asian center of biodiversity This has led to the suggestion that Taiwan may (Fujiwara et al. 2000). These reefs are known for have served as a“stepping-stone”for the north- their flourishing communities (Wells ward and eastward dispersal of shallow reef 1988). The Taiwan Strait, situated to the west organisms (Dai 1991a). between Taiwan and mainland China, is a shallow The octocoral fauna of Taiwan has been the channel, as opposed to the eastern region where subject of several studies. An early collection of the submarine topology drops steeply close to the soft corals conducted by Utinomi in 1938, resulted nearby Ryukyu Archipelago (Chen 1999). The in the description of 3 new species: geographic setting of Taiwan, between the West racemosa, formosana, and Asterospi- Pacific Ocean and East China Sea and at the cularia laurae (Utinomi 1950a b 1951). This col-

*To whom correspondence and reprint requests should be addressed. Tel: 972-3-6409090. Fax: 972-3-6409403. E-mail: [email protected] 548 Benayahu et al. -- Living Soft Corals of Southern Taiwan 549 lection was carried out mainly along the southern MATERIALS AND METHODS coast of Taiwan, and yielded 21 octocoral species of the families Clavulariidae, Tubiporidae, Collecting trips were conducted in 2 regions Xeniidae, Asterospiculariidae, Alcyoniidae, (Fig. 1): Kenting National Park, Nanwan Bay in Nephtheidae, Helioporidae, and Briareidae southern Taiwan (21 55'N; 120 45'E) during (Utinomi 1959). Later, Verseveldt (1982a) June-July 1994, Nov. °1998, and Mar.° 2004; and described an additional new species, Siphono- the offshore Green I. (Lutao in Chinese), southeast gorgia lobata (family Nidaliidae), from Taiwan. The of Taiwan (22 40'N; 121 29'E) during June-July 18 species of gorgonians listed for southern 1994. In Nanwan° Bay, collections° were made on Taiwan, all of which were found to be new records the fringing reefs including Banana Bay and for the region (Chen and Chang 1991), are like- Wanitun, mostly to a maximal depth of 12 m, and wise notable, but are beyond the scope of the cur- on the submerged offshore Sea Mount reef, down rent paper. Fujiwara et al. (2000) reported that to 31 m. At Green I., collections were made along species diversity of the Taiwanese reefs is relative- the western and southwestern sides down to a ly high, with 70 soft-coral and gorgonian species. depth of 31 m. The reefs were mostly reached by Dai (1991a b) demonstrated that soft corals in boat, and careful examination of a variety of niches Taiwan are abundant in the south, mainly on the in the different sites was carried out by scuba div- western side of Nanwan Bay. The reefs of Kenting ing. Approximately 230 samples were collected, National Park in southern Taiwan are very rich, encompassing the variety of species found on the with extensively developed soft-coral communities. reefs. Underwater abundance estimates of the They dominate the benthos, comprising in some genera were made visually and divided into 4 cate- sites up to 70% of the total coral cover (Dai 1993). gories: rare, sporadic, abundant, and dominant In these reefs, species of the genera , (see also Benayahu 1995). Prior to collection, , and are highly abundant most of the colonies were in situ photographed, and found even in habitats exposed both to strong using a Nikonos V camera with a close-up attach- tidal currents and typhoons (Dai 1988 1991b). A ment. Samples were fixed in 4% formalin in sea- checklist of the soft corals from southern Taiwan (Dai 1991a) included 42 species and an additional 7 species that remained unassigned. Among the 26°N N identified species of this list, Utinomi (1959) had already reported 13 species, and therefore 29 species appeared to be new records for the region. At present, there is a gap in the knowledge of soft corals from other regions of Taiwan. 25°N Intrigued by the early taxonomic reports on the soft corals of Taiwan (Utinomi 1950a b 1951 1959), their high abundance, and ecological impor- tance (Dai 1988 1990 1991a b 1993; Fujiwara et 24°N al. 2000), we conducted a comprehensive soft- Penghu Is. TAIWAN coral survey there. The present paper reports on reef sites located in both the East China Sea and the Pacific side of southern Taiwan. It provides a 23 N systematic list of soft-coral species, as well as ° Green I. information on their abundance and distributional Green I. patterns on the reefs. The coerulea is abundant in some of the surveyed Lanyu sites, and therefore it is also included in the study. 22°N Since the coral reefs of Taiwan and Japan are closely linked, we also compared the studied soft- coral fauna to those of reefs of the adjacent south- ern Ryukyu Archipelago and assessed their bio- Nanwan geographic affinities. 21 N ° 120°E 121°E Fig. 1. Map of Taiwan with study sites indicated by asterisks. 550 Zoological Studies 43(3): 548-560 (2004) water, rinsed in fresh water after 24 h, and then Zoology, Tel Aviv University, Israel. Some mem- transferred to 70% ethyl alcohol. Sclerites were bers of the families Alcyoniidea, Nephtheidae, obtained by dissolving the tissues in 10% sodium Nidaliidae, and Xeniidae are still being examined. hypochlorite. Identification of the Alcyoniidae species of the collection was in great part facilitated by compar- RESULTS isons with permanent sclerite preparations from type material kept in the Zoological Museum, The examined material yielded 69 species, of Department of Zoology, Tel Aviv University, Israel which 1 is a new species (Benayahu and Perkol- (ZMTAU) and at the Nationaal Natuurhistorisch Finkel 2004) and 43 are new zoogeographical Museum, formerly Rijksmuseum van Natuurlijke records for Taiwan (Table 1). Underwater pho- Historie, Leiden, The Netherlands. All material is tographs of some of the soft corals collected in the housed in the Zoological Museum, Department of current study are presented as indicated below.

Table 1. List of the species of Octocorallia of the orders Bock, 1938 and Alcyonacea Lamouroux, 1816 from the reefs of Taiwan with indication of Museum inventory numbers (ZMTAU Co) and previous references. NR, new record; 1, Utinomi 1959; 2, Dai 1991a; * occurrence of members of the family Alcyoniidae in the southern Ryukyu Archipelago, Japan following Benayahu 1995 2002; ** see Benayahu and Perkol-Finkel 2004

Classification References Classification References

Helioporidae Blainville, 1830 30252 Heliopora Moseley, 1786 Lobophytum sarcophytoides Moser, 1919 2 Heliopora coerulea (Pallas, 1766) 1, 2 ZMTAU Co 30239, 30250 ZMTAU Co 30273 Lobophytum venustum Tixier-Durivault, 1957* NR Clavulariidae Hickson, 1894 ZMTAU Co 29161, 29163, 29179, 30237, Genus Clavularia Blainville, 1830 30248 Clavularia inflata Schenk, 1896 1 Genus Paraminabea (Williams & Alderslade, 1999) ZMTAU Co 29197 Paraminabea aldersladei (Williams, 1992) NR Tubiporidae Ehrenberg, 1828 ZMTAU Co 29184, 29185, 30278, 30289 Genus Linnaeus, 1758 Genus Alderslade, 2000 Tubipora musica Linnaeus, 1758 1, 2 fulvum (Forskål, 1775)* NR ZMTAU Co 29193, 30169, 30256 ZMTAU Co 29200 Alcyoniidae Lamouroux, 1812 Genus Sarcophyton Lesson, 1834 Genus Gray, 1869 Sarcophyton buitendijki Verseveldt, 1982 NR (Macfadyen, 1936)* NR ZMTAU Co 29169 ZMTAU Co 28932, 28948, 28949, 30186, Sarcophyton crassocaule Moser, 1919* 2 30188, 30220, 30223 ZMTAU Co 29173, 30254 Cladiella daphnae Ofwegen & Benayahu, 1992 NR Sarcophyton ehrenbergi von Marenzeller, 1, 2 ZMTAU Co 30258 1886* Cladiella pachyclados (Klunzinger, 1877)* 1, 2 ZMTAU Co 30241, 30261 ZMTAU Co 30201 (Quoy & Gaimard, 1833)* 1, 2 Cladiella tuberculosa (Quoy & Gaimard, 1833) NR ZMTAU Co 29170, 29172, 29174, 29180, ZMTAU Co 29158, 30240, 30260, 30267 30238, 30242, 30243, 30244, 30246, 30251, Genus Putter, 1990 30268 Eleutherobia grayi (Thomson & Dean, 1931)* NR Sarcophyton infundibuliforme Tixier-Durivault, NR ZMTAU Co 30271 1958* Genus Alderslade, 2000 ZMTAU Co 29177, 30255 Klyxum simplex (Thomson & Dean, 1931)* 2 Sarcophyton nanwanensis Benayahu & ** ZMTAU Co 30195, 30219, 30245 Perkol-Finkel, 2004 Genus Lobophytum von Marenzeller, 1886 ZMTAU Co 30253 Lobophytum crassum von Marenzeller, 1886* 2 Sarcophyton tenuispiculatum Thomson & NR ZMTAU Co 28924, 29159, 29160, 29162, Dean, 1931* 29168 ZMTAU Co 29175 Lobophytum pauciflorum (Ehrenberg, 1834)* 2 Sarcophyton trocheliophorum von Marenzeller, 1, 2 ZMTAU Co 29164, 29165, 29166, 29167, 1886* Benayahu et al. -- Living Soft Corals of Southern Taiwan 551

Table 1. (Cont.)

Classification References Classification References

ZMTAU Co 29171, 29176, 29178, 30247 30218 Genus Sinularia May, 1898 Sinularia rigida (Dana, 1846) NR Sinularia brassica May, 1898* NR ZMTAU Co 28934 ZMTAU Co 28926, 30193, 30233 Sinularia triangula Tixier-Durivault, 1970 NR Sinularia compacta Tixier-Durivault, 1970 NR ZMTAU Co 30203, 30337 ZMTAU Co 30229 Sinularia variabilis Tixier-Durivault, 1945* NR Sinularia compressa Tixier-Durivault, 1945* NR ZMTAU Co 28942, 30204, 30182, 30215, ZMTAU Co 28940 30222, 30231 Sinularia discrepans Tixier-Durivault, 1970 NR Nephtheidae Gray, 1862 ZMTAU Co 28941 Genus Gray, 1869 Sinularia dissecta Tixier-Durivault, 1945 NR Capnella fungiformis Kükenthal, 1904 NR ZMTAU Co 30236 ZMTAU Co 28911, 28893, 28894 Sinularia erecta Tixier-Durivault, 1945* NR Genus Lemnalia Gray, 1868 ZMTAU Co 30187, 30208, 30225 Lemnalia flava (May, 1898) NR Sinularia exilis Tixier-Durivault, 1970 2 ZMTAU Co 28909 ZMTAU Co 30217 Genus Kükenthal, 1913 Sinularia facile Tixier-Durivault, 1970 2 Paralemnalia thyrsoides (Ehrenberg, 1834) 1, 2 ZMTAU Co 30200 ZMTAU Co 28905, 28906 (Quoy & Gaimard, 1833)* 2 Genus Studer, 1887 ZMTAU Co 28923, 28927, 30191, 30209, Scleronephthya gracillimum (Kükenthal, 1906) NR 30262 ZMTAU Co 29196, 29199 Sinularia gibberosa Tixier-Durivault, 1970* 2 Xeniidae Ehrenberg, 1828 ZMTAU Co 28930, 28938, 28939, 30190 Genus Anthelia Lamarck, 1816 Sinularia granosa Tixier-Durivault, 1970 2 Anthelia glauca Lamarck, 1816 NR ZMTAU Co 30192, 30214, 30285 ZMTAU Co 28883, 28888 Sinularia grayi Tixier-Durivault, 1945 NR Genus Asterospicularia Utinomi, 1951 ZMTAU Co 28925 Asterospicularia laurae Utinomi, 1951 Sinularia higai Benayahu, 2002* NR ZMTAU Co 32318 ZMTAU Co 30227 Genus Cespitularia Milne-Edwards, 1857 Sinularia humesi Verseveldt, 1968* NR Cespitularia caerulea May, 1898 NR ZMTAU Co 30206 ZMTAU Co 28886, 28877, 28880 Sinularia intacta Tixier-Durivault, 1970 NR Cespitularia Gohar, 1938 1, 2 ZMTAU Co 30216 ZMTAU Co 30277 Sinularia leptoclados (Ehrenberg, 1834)* NR Genus Kölliker, 1874 ZMTAU Co 28922, 28933, 28936, 28947, Heteroxenia medioensis Roxas, 1933 NR 30198, 30189, 30205, 30210, 30221, 30228, ZMTAU Co 30266 30234, 30235 Heteroxenia pinnata Roxas, 1933 NR Sinularia lochmodes Kolonko, 1926* 2 ZMTAU Co 28878 ZMTAU Co 28944, 28945, 30194, 30196 Genus Sansibia Aldersalde, 2000 Sinularia manaarensis Verseveldt, 1980 NR Sansibia flava (May, 1899) NR ZMTAU Co 28935, 30212 ZMTAU Co 28881, 28884, 28892, 30269 Sinularia maxima Verseveldt, 1971 NR Genus Lammarck, 1816 ZMTAU Co 30232 Xenia hicksoni Ashworth, 1899 NR Sinularia mollis Kolonko, 1926* 2 ZMTAU Co 28890 ZMTAU Co 28931, 28946 Xenia kükenthali Roxas, 1993 NR Sinularia nanolobata Verseveldt, 1977 NR ZMTAU Co 28889, 28891 ZMTAU Co 30224 Xenia lillieae Roxas, 1933 NR Sinularia notanda Tixier-Durivault, 1966 NR ZMTAU Co 30259 ZMTAU Co 28928, 28929, 30213 Xenia puerto-galerae Roxas, 1933 NR Sinularia numerosa Tixier-Durivault, 1970* 2 ZMTAU Co 28882 ZMTAU Co 30197, 30202, 30226, 30230 Briareidae Blainville, 1830 Sinularia ovispiculata Tixier-Durivault, 1970* NR Genus Blainville, 1830 ZMTAU Co 28943 Briareum excavatum (Nutting, 1911) NR Sinularia polydactyla (Ehrenberg, 1834)* 1, 2 ZMTAU Co 28784, 29194, 29195, 30249, ZMTAU Co 30335 30265, 30283 Sinularia querciformis (Pratt, 1903)* NR Briareum violacea (Quoy and Gaimard, 1883) 1 ZMTAU Co 28913, 28919, 30199, 30207, ZMTAU Co 28724, 30272, 30274, 30279 552 Zoological Studies 43(3): 548-560 (2004)

There are remarkable differences between flexibilis (Fig. 8) (see also Ofwegen 2002). In addi- the soft-coral diversity and generic composition of tion, other widespread alcyoniid species known in Nanwan Bay and Green I., with the former being other Indo-Pacific reefs were found there, i.e., richer (Table 2, 57 and 19 species, respectively). Cladiella australis (Fig. 9), Klyxum simplex (Fig. Although several species were recorded in both 10), Lobophytum crassum, (Fig. 11) L. pauciflorum areas, such as Lobophytum venustum, (Fig. 12), Sarcophyton ehrenbergi (Fig. 13), S. Paraminabea aldersladei, Sarcophyton crasso- glaucum (Fig. 14), S. infundibuliforme (Fig. 15), caule (Fig. 2), Sansibia flava, and Briareum exca- and Sinularia variabilis (see also Benayahu 2002). vatum (Fig. 3), each region appears to have its Green I. stands in stark contrast, with the complete own unique faunistic components (Table 2). absence of any species of the genus Sinularia. Species of the family Alcyoniidae are dominant This finding, along with the absence of Cladiella inhabitants of Nanwan Bay (Figs. 4, 5), comprising and Klyxum species and the lower number of 88% of the total number of species there. The Lobophytum and Sarcophyton species around genus Sinularia is very common there, represent- Green I., primarily accounts for the lower number ed by 30 species including some of the most-wide- of species there. Soft corals of the family Xeniidae spread species of the genus, i.e., S. leptoclados constitute the main soft corals on the reefs there, (Fig. 6), S. polydactyla (Fig. 7), S. brassica, and S. where they are represented by a variety of species (Table 1) such as Cespitularia caerulea (Fig. 16), Xenia kükenthali, X. puerto-galerae, and Anthelia Table 2. Generic affiliation and abundance esti- glauca (Fig. 17). The xeniid assemblages are mates of soft-coral genera of Nanwan Bay and abundant on the reefs of the western and south- Green I. Taiwan western sides of Green I., where they form a high living coverage at a depth range of 3~10 m (Fig. Taxa Number of species Abundance 18). Other soft corals are found intermingled with- in the Xeniidae, including Clavularia inflata (Fig. Nanwan Bay Green I. Nanwan Bay Green I. 19), Capnella fungiformis (Fig. 20), Lemnalia flava Family Helioporidae (Fig. 21), and Paralemnalia thyrsoides. None of Heliopora 1 - Sporadic - the 4 latter species was found in Nanwan Bay. In Family Clavulariidae contrast, species of the family Xeniidae in Nanwan Clavularia - 1 - Rare Bay were observed only in shallow reef habitats, Family Tubiporidae as they also were in the tidal pools in Banana Bay. Tubipora 1 1 Rare Rare Interestingly, Sansibia flava was sporadically col- Family Alcyoniidae lected in both study regions. Asterospicularia lau- Cladiella 4 - Sporadic - rae (Fig. 22) also of the family Xeniidae (see Eleutherobia 1 - Rare - Alderslade 2001) has rarely been found in Nanwan Klyxum 1 - Sporadic - Bay. Lobophytum 4 1 Abundant Rare Paraminabea 1 1 Rare Rare The shallow fringing reefs of Nanwan Bay Rhytisma - 1 - Rare (3~12 m) are densely inhabited mainly by species Sarcophyton 7 2 Abundant Sporadic of the family Alcyoniidae in the genera Cladiella, Sinularia 30 - Dominant - Klyxum, Lobophytum, Sarcophyton, and Sinularia, Family Nephtheidae (Figs. 4, 5). They form large monospecific aggre- Capnella - 1 - Rare gations that monopolize the substrate and occa- Lemnalia - 1 - Rare sionally even become the most-dominant benthic Paralemnalia - 1 - Sporadic component on the reefs (see“Discussion”). In Scleronephthya - 1 Sporadic contrast, the deep reef habitat of Nanwan Bay (at Family Xeniidae “Sea Mount”) is characterized by sporadic Anthelia - 1 - Rare colonies of the Alcyoniidae, mainly in the genera Asterospicularia 1 - Rare - Cespitularia 1 1 Rare Dominant Lobophytum and Sarcophyton (Fig. 23). They are Heteroxenia 1 1 Rare Rare also smaller in size compared to those of the shal- Sansibia 1 1 Sporadic Sporadic lower reef (e.g., Figs. 4, 7, 15). They become Xenia 1 3 Rare Dominant scarcer below 18~20 m, where we mostly record- Family Briareidae ed assemblages of azooxanthellate species of the Briareum 2 1 Sporadic Sporadic genera (Fig. 24), Stereo- Total 57 19 nephthya, Umbellulifera (family Nephtheidae), and Benayahu et al. -- Living Soft Corals of Southern Taiwan 553

Chironephthya (Fig. 25) and (family Heteroxenia elisabethae, Sinularia mayi, and Nidaliidae). Identification to species level of these Cladiella sphaerophora; but excluding Nephthea specimens awaits future studies, and therefore erecta, the genus of which is not included in the they do not appear in Table 2. The azooxanthel- present study). Sixteen species listed by Dai late species Eleutherobia grayi (Fig. 26) and (1991a) were not recorded in the current survey Paraminabea aldersladei (family Alcyoniidae) were (i.e., Cladiella sphaerophora, Klyxum molle, K. found in both study regions, mostly growing on rotundum, Lobophtum batarum, L. mirabile, L. reef overhangs. At Green I., aggregations of the solidum, Sarcophyton stellatum, S. tortuosum, azooxanthellate species Scleronephthya gracilli- Sinularia densa, S. grandilobata, S. halversoni, S. mum (Fig. 27) are abundant along a wide depth inexplicita, S. mayi, S. muralis, S. scabra, and gradient, mainly in niches that lack direct sun radi- Heteroxenia elisabethae, as emended following ation, such as reef overhangs (Fig. 28), or beneath Alderslade 2000). Three of these species appear , colonies of branching stony corals (Fig. 29). in Utinomi s (1959) list but were not found in the Despite the differences in the faunistic fea- present survey. Absence of previously recorded tures of soft corals between Nanwan Bay and species (Utinomi 1959, Dai 1991a) in our survey Green I. (Table 2), in both regions they constitute may reflect temporal changes in soft-coral species an important faunistic component that significantly composition due to natural or man-made perturba- contributes to the living coverage. tions occurring on Taiwanese reefs (see below). Nonetheless, our findings provide a firm indication of the high soft-coral diversity on Taiwanese reefs. DISCUSSION Species of the family Alcyoniidae are domi- nant in Nanwan Bay, with a remarkably high diver- The overall objective of the present survey sity of Sinularia (Table 2). The 30 species of this was to investigate the soft-coral fauna of southern genus recorded in our survey place the Taiwanese Taiwan. The results indicate that soft corals are reefs among other Indo-Pacific sites known for abundant in both Nanwan Bay (East China Sea) their high scores of Sinularia species (Ofwegen and around Green I. (Pacific Ocean), totaling 69 2002). Our field observations confirmed previous species of the families Helioporidae, Clavulariidae, findings (Dai 1991a b 1993) that on certain reefs in Tubiporidae, Alcyoniidae, Nephtheidae, Xeniidae, this region, species assemblages of this family and Briareidae (Tables 1, 2). Among the 22 listed form high living coverage of up to 60%~70% of the genera of these families, the survey records 7 for total surface area. They often propagate by colony the 1st time from Taiwanese reefs, i.e., fission (Figs. 30, 31) and consequently form large Eleutherobia, Paraminabea, Rhytisma, Capnella, monospecific aggregations on reefs. Contrasting Lemnalia, Scleronephthya, and Sansibia. The cur- abundance patterns of the Xeniidae in Nanwan rent survey also yielded a new species Bay and Green I. probably reflect different envi- (Sarcopyhton nanwanensis Benayahu and Perkol- ronmental regimes prevailing in each of these 2 Finkel, 2004) and 43 new zoogeographical regions. Nanwan Bay suffers from occasional records, which constitute 63% of the total number typhoons in summer and autumn, which the rigid of species identified in the present study. colonies of the family Alcyoniidae are able to with- The present collection shares 25 species with stand (see also Dai 1988 1993). Even after severe previous studies by Utinomi (1959) and Dai storms, the remnants of these colonies undergo (1991a). Noteworthy is the recent collection (Mar. rapid regeneration, which enables them to effec- 2004) of Asterospicularia laurae (Table 1), whose tively reoccupy space, and may account for their type locality is the southern tip of Taiwan (Utinomi dominance on storm-swept reefs (Dai 1991b). 1951). The family Asterospiculariidae has recently Species of the family Xeniidae tend to prefer more- been abandoned, and the genus Asterospicularia protected reefs and calmer habitats than other has been transferred to the Xeniidae (Alderslade soft-coral species (Y.B., pers. observ.). Due to 2001). Emendations following the taxonomic revi- their fast asexual propagation by runner-like sion by Verseveldt (1982b) of Sarcophyton and stolons (Fig. 32), they can successfully exploit reef assessments by Fabricius and Alderslade (2001) substrate, and consequently even exclude other assigning Pachycalvularia to Briareum, reduce to 5 reef dwellers (e.g., Benayahu and Loya 1987, , the number of species from Utinomi s (1959) list Karlson et al. 1996). Interestingly, southern that were not found in the present collection (i.e., Taiwan and Green I. possess similar stony-coral Clavularia racemosa, Anthelia formosana, distributions of both and Faviidae 554 Zoological Studies 43(3): 548-560 (2004) species (Chen 1999) and of reef fishes (Shao et al. the Alcyoniidae are found in common in both 1997), suggesting that soft corals of the families Taiwan and the Ryukyu Archipelago. It is also Alcyoniidae and Xeniidae are more sensitive to the interesting to note that Sinularia higai, which has differential environmental regime around Taiwan recently been described from the southern Ryukyu compared to other taxa. Archipelago (Benayahu 2002), was also recorded To date, among the soft corals, species of the in this study (Table 1, Fig. 33). Furthermore, even family Alcyoniidae are undoubtedly the best known Sarcophytom nanwanensis which was recently throughout the Indo-Pacific region, mainly due to described from southern Taiwan, is also found in the taxonomic revisions of the 3 most common Okinawa (Benayahu and Perkol-Finkel 2004). It is genera of this family, Sinularia, Sarcophyton, and anticipated that similar patterns also exist for the 2 Lobophytum (Verseveldt 1980 1982b 1983). other important soft-coral families, Nephtheidae These revisions have contributed to increased and Xeniidae. A biogeographic analysis of soft knowledge of the species of the respective genera corals in the West Pacific Ocean and East China and their distributions among Indo-Pacific reefs, Sea should undoubtedly also include the particularly of Sinularia, the largest shallow-water Philippines (see also Chen 1999). Only a few soft-coral genus (Ofwegen 2002 and references papers have been published on the soft corals of , therein). The coral reefs of Taiwan and Japan are the latter s reefs (Ofwegen 2002), and therefore at closely linked by the northward flowing Kuroshio this stage, no conclusions can be made concern- Current, which brings warm water and larvae from ing their real diversity or on the distributional pat- reefs of the South China Sea and the Philippines terns of the soft-coral fauna throughout the entire (Fujiwara et al. 2000). Therefore, it is not surpris- Philippine-Japan I. Arc. It is worthy of note that ing that Imahara (1996) established a joint octoco- Cladiella daphnae was recorded in southern ral list for Japan and the adjacent seas including Taiwan (Table 1). This is the 1st record of this those of Taiwan. Data obtained on the Alcyoniidae species away from Tanzania, which is the species soft corals from southern Taiwan (this study) and type locality (Ofwegen and Benayahu 1992). Such from the southern Ryukyu Archipelago, Japan a distributional pattern of a species, from both (Benayahu 1995, 2002) enabled us to compare the sides of the Indo-Pacific region, is an indication of generic affiliations and abundance estimates within the need for further soft-coral surveys that will this family in the 2 regions (Table 3). In both determine the true range of distribution of various regions, the Alcyoniidae constitute the most-domi- soft coral taxa. nant soft-coral component. There is a close The coral reefs of Taiwan increasingly suffer resemblance between their respective faunas in from man-made perturbations due to coastal con- terms of generic composition, abundance, and struction, dredging, destructive fishing practices, number of species. Furthermore, 30 species of coral collection, unregulated tourist activities, and anchoring (Dai 1997, Fujiwara et al. 2000, Dai et Table 3. Number of species of the family al. 2002). In addition, pollution of various types, Alcyoniidae in Taiwan (this study) and the south- primarily derived from a nuclear power plant in ern Ryukyu Archipelago (Benayahu 1995 2002) Nanwan Bay, has been considered a potential threat to the well-being of the adjacent reefs (Jan et al. 2001, Dai et al. 2002). Some of the reefs of Taxa Number of species southern Taiwan suffered from extensive bleaching Taiwan Southern Ryukyu Archipelago during 1998 (Fig. 34), at which time coral cover was substantially reduced (Fujiwara et al. 2000). Family Alcyoniidae Cladiella 44The likelihood of devastating effects occurring as a Eleutherobia 11result of the above-mentioned factors threatens Klyxum 12the biodiversity of Taiwanese reefs, and strongly Lobophytum 48necessitates the promotion of conservation and Paraminabea 1-sustainable use of the reef resources there. The Protodendron -1current high diversity of the soft corals found in this Rhytisma 11survey further reinforces the need to implement Sarcophyton 79effective conservation policies and management Sinularia 30 25 programs in order to promote maintenance and Total 49 51 preservation for this unique reef ecosystem. Benayahu et al. -- Living Soft Corals of Southern Taiwan 555

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Fig. 2. Sarcophyton crassocaule Moser, 1919. Fig. 3. Briareum excavatum (Nutting, 1911). Fig. 4. Reef area in Nanwan Bay covered by Sinularia and Lobophytum. Fig. 5. Reef area in Nanwan Bay covered by Sarcophyton and Sinularia. Fig. 6. Sinularia leptoclados (Ehrenberg, 1834). Fig. 7. Sinularia polydactyla (Ehrenberg, 1834). Fig. 8. Aggregations of Sinularia flexibilis (Quoy and Gaimard, 1833). Fig. 9. Cladiella australis (Macfadyen, 1936). 556 Zoological Studies 43(3): 548-560 (2004)

10 11

12 13

14 15

16 17

Fig. 10. Klyxum simplex (Thomson and Dean, 1931). Fig. 11. Lobophytum crassum von Marenzeller, 1886. Fig. 12. Lobophytum pauciflorum (Ehrenberg, 1834). Fig. 13. Sarcophyton ehrenbergi von Marenzeller, 1886. Fig. 14. Sarcophyton glaucum (Quoy and Gaimard, 1833). Fig. 15. Sarcophyton infundibuliforme Tixier-Durivault, 1958. Fig. 16. Cespitularia caerulea May, 1898. Fig. 17. Anthelia glauca Lamarck, 1816. Benayahu et al. -- Living Soft Corals of Southern Taiwan 557

18 19

20 21

22 23

24 25

Fig. 18. Assemblages of Xeniidae on reefs on the western side of Green I. Fig. 19. Clavularia inflata Schenk, 1896. Fig. 20. Capnella fungiformis Kükenthal, 1904. Fig. 21. Lemnalia flava (May, 1898). Fig. 22. Asterospicularia laurae Utinomi, 1951. Fig. 23. Deep reef habitat of Nanwan Bay (at“Sea Mount”) with colonies of Alcyoniidae. Fig. 24. Dendronephthya sp. from the deep reef of Nanwan Bay. Fig. 25. Chironephthya sp. from the deep reef of Nanwan Bay. 558 Zoological Studies 43(3): 548-560 (2004)

26 27

28 29

30 31

32 33

Fig. 26. Eleutherobia grayi (Thomson and Dean, 1931) with Fig. 27. Scleronephthya gracillimum (Kükenthal, 1906) with polyps expanded. polyps expanded. Fig. 28. Aggregations of Scleronephthya gracillimum (Kükenthal, Fig. 29. Aggregations of Scleronephthya gracillimum (Kükenthal, 1906) growing on a reef overhang. 1906) growing beneath colonies of the stony coral Fig. 30. Fission of a Sinularia colony. Acropora. Fig. 32. Asexual propagation by runner-like stolons of Fig. 31. Fission of a Sinularia colony. Cespitularia. Fig. 33. Sinularia higai Benayahu, 2002. Benayahu et al. -- Living Soft Corals of Southern Taiwan 559

indicating a pattern congruent with sea surface tempera- 34 tures and currents: examples from Acropora and Faviidae corals. Zool. Stud. 38: 119-129. Chen CC, KH Chang. 1991. Gorgonacea (Coelenterata: : Octocorallia) of southern Taiwan. Bull. Inst. Zool. Acad. Sinica 30: 149-182. Dai CF. 1988. Coral communities of southern Taiwan. Proc. 6th Int. Symp. Townsville, Queensland, Australia. 2: 647-652. Dai CF. 1990. Interspecific competition in Taiwanese corals with special reference to interactions between alcy- onacean and scleractinian corals. Mar. Ecol.-Prog. Ser. 60: 291-297. Dai CF. 1991a. Reef environment and coral fauna of southern Taiwan. Res. Bull. 354: 1-28. Fig. 34. Bleached colony of Sinularia sp. in Nanwan Bay. Dai CF. 1991b. Distribution and adaptive strategies of alcy- onacean corals in Nanwan Bay, Taiwan. Hydrobiologia 216: 241-246. Dai CF. 1993. Patterns of coral distribution and benthic space Acknowledgments: This study was made possi- partitioning on the fringing reefs of southern Taiwan. Mar. ble due to a grant from the National Science Ecol.- P.S.Z.N.I. 14: 185-204. Council, R.O.C. that allowed Y.B. to visit Taiwan. Dai CF. 1997. Assessment of the present health of coral reefs We would like to express our gratitude for the in Taiwan. In RW Grigg, C Birkeland, eds. Status of coral enthusiastic assistance of K. Soong and Y. Tsay reefs in the Pacific. Manoa, Hawaii: Sea Grant Program, Univ. of Hawaii, pp. 123-131. during the field trips. We are also grateful for the Dai CF, G Chen, M Inaba, K Iwao, F Iwase, Y Kakuma, K highly constructive critical comments of L. P. van Kajiwara, T Kimura, K Nomura, K Oki, K Sakai, T Ofwegen during part of the identification work and Shibuno, H Yamano, M Yoshida. 2002. Status of coral regarding the manuscript. Y.B. thanks the late J. reefs in East Asia: China, Japan, Korea and Taiwan. 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